Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain

ANGELICA TORICES, PHILIP J. CURRIE, JOSE IGNACIO CANUDO, and XABIER PEREDA-SUBERBIOLA

Torices, A., Currie, P.J., Canudo, J.I., and Pereda-Suberbiola, X. 2015. Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Palaeontologica Polonica 60 (3): 611–626.

The dinosaur record in the South Pyrenees Basin is diverse and rich. A total of 142 theropod teeth were studied for this paper, which constitutes one of the richest samples for these remains in Europe. Eight upper Campanian to upper Maastrichtian outcrops from the Pyrenees produced six non-avian theropod taxa (Theropoda indet., Coelurosauria indet., ?Richardoestesia, ?Dromaeosauridae indet., ?Pyroraptor olympius, ?Paronychodon). These six taxa are added to two previously described theropods (a Richardoestesia-like form and a possible ornithomimosaurid), indicating that there was considerable theropod diversity on the Iberian Peninsula during the Late Cretaceous.

Key words: Dinosauria, Theropoda, teeth, Cretaceous, Spain, South Pyrenees.

Angelica Torices [[email protected]] and Philip J. Currie [[email protected]], Department of Biological Sciences, University of Alberta, CW 405 Biological Sciences Centre, Edmonton, Alberta, Canada. José Ignacio Canudo [[email protected]], Grupo Aragosaurus-IUCA, Paleontología, Facultad de Ciencias, Univer- sidad de Zaragoza, Pedro Cerbuna 12, 50009 Zaragoza, Spain. Xabier Pereda-Suberbiola [[email protected]], Universidad del País Vasco/Euskal Herriko Unibertsitatea (UPV/ EHU), Facultad de Ciencia y Tecnología, Dpto. Estratigrafía y Paleontología, Apdo. 644, 48080 Bilbao, Spain.

Received 12 October 2012, accepted 28 October 2013, available online 30 October 2013.

Copyright © 2015 A. Torices et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

the rest of Spain (Pol et al. 1992; Pereda-Suberbiola 1999b, Introduction Prieto-Márquez et al. 2000; Company 2004). On the Iberian Peninsula, few theropod teeth have been Institutional abbreviations.—AMNH, American Natural recovered in association with their cranial and postcranial History Museum, New York, USA; DPM, Departamento de skeletal remains. In southern Europe, isolated theropod teeth Paleontología de Madrid, Complutense University of Ma- tend to be scattered amongst the remains of other dinosaurs, drid, Spain; MCNA, Museo de Ciencias Naturales de Álava, and have no relation to skeletal remains attributable to any Vitoria-Gasteiz, Spain; MPZ, Museo Paleontológico, Uni- defined theropod taxon (Buffetaut and Le Loeuff 1991). Py- versidad de Zaragoza, Zaragoza, Spain. roraptor olympius is an exception in that two isolated teeth were associated with postcranial skeletal remains (Allain and Other abbreviations.—FABL, fore-aft-basal length; PCA, Taquet 2000). The vast majority of Upper Cretaceous (Cam- principal components analysis. panian–Maastrichtian) theropod fossils in Europe consist of isolated teeth, and they provide the greatest potential for identifying which theropod taxa were present in the region. Material and methods The dinosaur record in the South Pyrenees Basin is rich, comprising 147 dinosaur sites that can be assigned to four Eight localities (from east to west: Laño in Treviño; Blasi 1, time intervals from late Campanian to late Maastrichtian 2B and 3 in Huesca; Vicari 4, Montrebei, Figuerola 2, and (López-Martínez 2003b). In this paper, teeth are used to Fontllonga 6 in Lleida) produced the 142 theropod teeth used identify the Upper Cretaceous theropod dinosaurs from this in this study. Laño, the richest site, produced 120 of these region. Laño is the richest site for theropod teeth in this area, teeth. The sample is relatively small, when compared with and for the Upper Cretaceous of Europe. Outside of the South Upper Cretaceous North American sites, each of which can Pyrenees Basin, fewer than a dozen teeth have been found in produce up to thousands of specimens (Currie et al. 1990;

Acta Palaeontol. Pol. 60 (3): 611–626, 2015 http://dx.doi.org/10.4202/app.2012.0121 612 ACTA PALAEONTOLOGICA POLONICA 60 (3), 2015

Farlow et al. 1991; Sankey 2001; Sankey et al. 2002; Sam- man et al. 2005; Smith 2005). However, for Europe, this is one of the largest Cretaceous theropod tooth samples that has been examined. Several publications (Csziki and Grig- orescu 1998; Codrea et al. 2002; Laurent et al. 2002; Allain and Pereda-Suberbiola 2003; Smith et al. 2005) mention European theropod teeth without providing figures on their abundances. Where numbers are given, the sample varies from one to 58 (Buffetaut et al. 1986, 1988; Antunes and Sigogneau-Russell 1991, 1992; Sigé et al. 1997; Le Loeuff and Buffetaut 1998; Allain and Taquet 2000; Garcia et al. 2000; Laurent 2002; Lindgren et al. 2009; Ösi et al. 2010). Theropod teeth were measured for this study using a Lei- ca Wild M-10 stereo microscope. Measurements included: crown height, fore-aft-basal length (FABL), basal width of the crown, anterior denticle density, and posterior denticle density. The last two measurements were recorded as the number of denticles per millimeter, and were taken at the mid-heights of the carinae. Qualitative characteristics (such as the shape of a denticle) were noted, as these are often important in the identification of theropod teeth (Currie et al. 1990). For example, Troodon teeth have strongly hooked denticles, velociraptorine teeth have distally-hooked and sharply-pointed denticles, and dromaeosaurines and other theropods have denticles that are squared-off and chisel-like distally. The presence of longitudinal ridges was noted, as this is common in teeth attributed to Paronychodon lacustris. Statistical analyses were carried out using SPSS 21.0 (IBM Corp. Released 2012. IBM SPSS Statistics for Win- dows, Version 21.0. Armonk, NY: IBM Corp.). A total of 650 teeth—representing 21 species and eleven families—were included in these analyses. Bivariate analyses of the Pyrene- an sample and taxa were compared with a sample of a further 17 theropod species, largely from North America, but with some South American (Aucasaurus, Mapusaurus) and Asian (Fukuiraptor, Tarbosaurus) species. These analyses were performed to compare the relationship between tooth size and denticles per millimeter. Principal components analysis (PCA) was performed to separate the specimens according to the variance of the height, FABL, basal width, and denticle Fig. 1. A. Locations of the palaeontological sites of Laño, Vicari 4, Mon- size. The data were log-transformed for the analyses. trebei, Fontllonga 6, Figuerola 2, and Blasi. B. Correlation of the upper- Three discriminant analyses were performed on the most Cretaceous and lowermost Tertiary deposits in the southern Pyrenees, log-transformed values of the Pyrenean sample to verify the showing the stratigraphic levels of the studied localities. MPU, Mid-Paleo- cene unconformity; S1, S2, depositional sequences (Robador 2005). preliminary assignments of the specimens. Analyses were carried out at both species and family level. pollen (Galbrun et al. 1993; López-Martínez et al. 1998; López-Martínez 2003a) and magnetostratigraphy (Galbrun et al. 1993; López-Martínez et al. 2006; Pereda-Suberbiola et al. Geological setting 2009; Riera et al. 2009). The first site, where practically all the material has been The Late Cretaceous theropod localities considered here are found, is Laño in the Basque-Cantabrian Region. The Laño all located in the South Pyrenees Basin (Garrido Mejías and site (Fig. 1) is situated in an abandoned sand quarry within Ríos Aragües 1972), extending over some 1500 km from east the Condado de Treviño, 25 km south of the city of Vitoria- (Provence, France) to west (Cantabrian Platform), with an av- Gasteiz (Alava province). This quarry is on the south flank of erage width of 200 km from north (Aquitaine Basin) to south the Sub-Cantabrian Synclinorium, a great structure composed (Ebro Basin). The eight sites have been dated by means of mostly of Upper Cretaceous and Palaeogene deposits that ex- biostratigraphy using planktonic foraminifera, charophytes, tend more than 100 km from east to west (Astibia et al. 1987, TORICES ET AL.—LATE CRETACEOUS THEROPOD DINOSAURS FROM SPAIN 613

1990; Baceta et al. 1999). Most of the vertebrate fossils have Blasisaurus in Blasi 1 (Cruzado-Caballero et al. 2010) and been recovered from upper Campanian–lower Maastrichtian Arenysaurus in Blasi 2 (Pereda-Suberbiola et al. 2009). siliciclastic rocks in three beds, known as L1A, L1B, and L2. The sedimentological interpretations of these formations These beds are part of a Late Cretaceous alluvial system com- and their relationships are complex. Several authors have sup- posed mainly of fluvial sand and silt, interpreted as channel ported a model of barrier islands and coastal lagoons for the deposits of an extensive braided river system (Astibia et al. formation of these deposits (Nagtegaal et al. 1983; Diaz-Mo- 1990). Unusually, the fossiliferous beds are associated with lina 1987; Diaz-Molina et al. 2007). According to this mod- iron crusts, and the fossil bones are often covered by iron el, longitudinal platform currents would have built a barrier oxides (Elorza et al. 1999; Pereda-Suberbiola et al. 2000). island that became the Arén Sandstone deposits. After that, Laño has yielded a diverse continental vertebrate assemblage, Tremp Formation deposits were formed in lagoons protected which consists of nearly 40 species of actinopterygians, lis- from the waves. At the top of the Arén Sandstone, palaeosols, samphibians, lepidosaurs, turtles, crocodyliforms, dinosaurs, ferruginous encrustations and nodules are commonly found, pterosaurs, and mammals. In addition to theropods, other which implies that the Tremp Formation marly sediments did dinosaurs include the titanosaurian sauropod Lirainosaurus not form until after a period of emersion and subaerial expo- astibiae (Sanz et al. 1999; Díez Díaz et al. 2011, 2012), the or- sure of the barrier island (Díaz-Molina 1987) (Fig. 1). nithopod Rhabdodon (Pereda-Suberbiola and Sanz 1999) and the ankylosaur Struthiosaurus (Pereda-Suberbiola 1999a). The seven other sites are in the Montsec Thrust sheet Systematic palaeontology (Tremp Syncline in the north, Tremp-Graus Basin) and the Sierras Marginales Thrust sheet (Ager Syncline in the south). Dinosauria Owen, 1842 The Montsec Thrust separates these two structures. In both Saurischia Seeley, 1888 synclines, the Cretaceous–Tertiary transition is represented by red-bed deposits of the Tremp Formation (Mey et al. Theropoda Marsh, 1881 1968; López-Martínez et al. 1998, 1999; Riera et al. 2009). Theropoda indet. Two of the sites are in the Ager Syncline at the base of Unit Fig. 2A, D–H. 2; Figuerola 2 is late Campanian in age, and Fontllonga 6 is Material.—There are thirteen large, isolated theropod teeth early Maastrichtian (López-Martínez et al. 1998). The rest of from Blasi 1, 2B, and 3, Laño, and Montrebei (upper Cam- the sites are in the Tremp Syncline. Vicari 4 and Montrebei panian–upper Maastrichtian) that can be separated into two are found at the base of the Tremp Formation near the contact morphotypes. Morphotype 1: two teeth (MPZ2004/3 and with the Arén Sandstone; the age of Vicari 4 is late Campa- 4/4) from Blasi 1, one tooth (MPZ2004/5) from Blasi 2B, and nian, and Montrebei is latest Campanian to basal Maastrich- two teeth (MPZ2004/8 and 98/67) from Blasi 3); four teeth tian (Vicens et al. 2001). Blasi 1 is in the Arén Formation, (MCNA 22051, 4520, 14521, and 1852) from Laño (lateral and Blasi 2B and 3 in the Tremp Formation; all three are late equivalent of Sedano Formation, upper Campanian–lower Maastrichtian (López-Martínez et al. 2001; López-Martínez Maastrichtian); one tooth (DPM-MON-T10) from Mon- 2003b; Pereda-Suberbiola et al. 2009; Fig. 1). trebei (Tremp Formation, upper Campanian–lower Maas- The Figuerola 2 fossiliferous level consists of 2.5 m of trichtian) (Fig. 2E–H; SOM: Table 1 [Suplementary Online grey marl, upon which is superimposed a layer of 0.5 m made Material available at http://app.pan.pl/SOM/app60-Torices_ up exclusively of oncolites. Above this layer there is 4.5 m of etal_SOM.pdf]). Morphotype 2: one tooth (MPZ98/68) from grey marl. In the marl, bone fragments, scales of bony fish, Blasi 1 and two teeth (MCNA 1853, 14522) from Laño (Fig. ray teeth, dinosaur eggshells, and theropod teeth have been 2A, D; SOM: Table 1). found. At the Fontllonga 6 site, theropod teeth appear in a level composed of 1.8 m of grey marl with yellowish and Description brownish alterations. Fossil remains are found covered with Morphotype 1.—Teeth have crowns whose height varies be- carbonate crusts and oncolitic layers. tween 18 and 62 mm, FABL between 11 and 27 mm, and Montrebei and Vicari 4 theropod teeth are found in thin basal widths between 4.4 and 13 mm. Their carinae have levels (0.5 m) of dark marl that is rich in organic matter asso- large rectangular denticles. Their anterior denticle densities ciated with fish teeth and scales, crocodile teeth, and eggshells. vary between 2 and 5 denticles/mm, and posterior denticle Blasi 2B occurs in a 6.5 m thick interval of grey marl that densities range between 2 and 3 denticles/mm. The largest overlie the sandstone of Blasi 1. Nearly 5000 kg of sediment anterior and posterior denticles are more or less the same have been washed, resulting in the extraction of vertebrate size, although the anterior ones tend to be a bit smaller than microfossils (fish, amphibians, squamates, and turtles), in- the posterior ones. The teeth are vertically elongate with cluding some dinosaur teeth (theropods, hadrosaurids) and straight or slightly curved borders in lateral view. eggshell fragments. Plant debris, charophytes and gastropods Variation in tooth morphology along the jaw has been ob- are common (López-Martínez et al. 2001; Blain et al. 2010). served in theropods and there is a tendancy towards increased Two hadrosaurids have been described in the Blasi section— labiolateral compression towards the posterior regions of the 614 ACTA PALAEONTOLOGICA POLONICA 60 (3), 2015

AB C D

EF G H

Fig. 2. Theropod dinosaurs teeth from upper Campanian–lower Maastrichtian, Laño. A, D. Theropoda indet. Morphotype 2. A. MCNA 14522. D. MCNA 1853. B, C. ?Pyroraptor olympius Allain and Taquet, 2000. B. MCNA 14623. C. MCNA 14624. E–H. Theropoda indet. Morphotype 1. E. MCNA 1852. F. MCNA 14520. G. MCNA 14521. H. MCNA 2205. All lateral views. Scale bars 5 mm. jaws (Currie et al. 1990; Smith 2005; Reichel 2010). Some The only species of large theropod defined for the Late Morphotype 1 teeth have rounded cross-sections that may Cretaceous of Europe is, at the moment, Tarascosaurus sal- indicate that they are from anterior positions in the jaws. luvicus from Var (southern France), which is represented by Morphotype 2.—Teeth are smaller (between 20 and 24 mm) two dorsal vertebrae and a femur (Le Loeuff and Buffetaut than those of Morphotype 1 (although there is slight overlap 1991). This taxon is probably an abelisauroid (Carrano and between the ranges) and are strongly curved. Their FABLs Sampson 2008) but it was considered a nomen dubium by are 12 and 13 mm, and their basal widths are 6.2 and 7.3 mm. Allain and Pereda-Suberbiola (2003). In Laño, this taxon Their denticles have the same morphologies as Morphotype was identified on the basis of two femora by Le Loeuff and 1. Anterior denticle density is 4 denticles/mm and posterior Buffetaut (1991) and Le Loeuff (1992), and afterwards it has denticle density is 3 denticles/mm. subsequently been referred to Neoceratosauria indet. (Pere- da-Suberbiola 1999b; Pereda-Suberbiola et al. 2000). Discussion Abelisauroidea, the clade to which Tarascosaurus be- Morphotype 1.—In Europe, several large theropods (includ- longs (Le Loeuff and Buffetaut 1991; Le Loeuff 1992), is ing abelisaurids and megalosaurids) have been described characteristic of the late Mesozoic of Gondwana (Sereno et from Upper Cretaceous sites (Le Loeuff and Buffetaut 1991; al. 2004); they could have reached Europe during the Early Le Loeuff 1992; Pereda-Suberbiola 1999b; Pereda-Suberbi- Cretaceous (Pereda-Suberbiola 2009) or even during the Ju- ola et al. 2000). Remains attributed to Megalosauridae (teeth rassic as recently suggested (Ezcurra and Agnolin 2012; see and a radius) have been described in Portugal and Spain, but Pol and Rauhut 2012; Rauhut 2012 for a different interpreta- this fragmentary material is probably not diagnostic enough tion). Camarillasaurus cirugedae (Barremian; Sánchez-Her- to classify at family level (Pereda-Suberbiola 1999b). nandez and Benton 2014), Genusaurus sisterornis (Albian; TORICES ET AL.—LATE CRETACEOUS THEROPOD DINOSAURS FROM SPAIN 615

Accarie et al. 1995), Tarascosaurus salluvicus (Campanian), terial is found, these teeth should more conservatively be Betasuchus bredai (Maastrichtian) and other taxa not formal- referred to as indeterminate Theropoda. ly named from the Campanian–Maastrichtian (Buffetaut and Le Loeuff 1991; Allain and Pereda-Suberbiola 2003) indi- Coelurosauria Huene, 1914 cate the presence of multiple ceratosaur lineages (including Coelurosauria indet. abelisauroids) in Europe through the end of the Cretaceous (Carrano and Sampson 2008). According to Allain (1998), Material.—Fifty isolated teeth: MPZ98/79–82 from Blasi European species could represent an independent line of cer- 2B (Tremp Formation, late Maastrichtian); MCNA 14523– atosaurs that evolved in isolation. 14565 from Laño (lateral equivalent of Sedano Formation, Neoceratosaurs are known from the Late Jurassic to the upper Campanian–lower Maastrichtian); DPM-MON-T3, latest Cretaceous (Carrano et al. 2002; Sereno et al. 2004; T6 from Montrebei (Tremp Formation, upper Campanian– Carrano and Sampson 2008) in other parts of the world, and lower Maastrichtian); DPM-VIR4-T5 from Vicari 4 (Tremp include both small sized theropods (such as Masiakasaurus, Formation, upper Campanian) (Fig. 3M–P; SOM: Table 1). which is 1.8–2 m in total length; Carrano et al. 2002) and large Description.—The indeterminate coelurosaurian teeth are ones (such as Carnotaurus at 7.5 m, Bonaparte et al. 1990; small and lack denticles. Height varies between 1.5 and and Majungatholus at 6–7 m, Krause et al. 2007). Allain et 8.1 mm. The anterior carina of each is convex, whereas the al. (2007) described the neoceratosaur Berbesaurus liassicus posterior one is concave in lateral view. from the Jurassic of Morocco, but not everyone agrees with The most prominent features of the indeterminate coe- its classification (Carrano and Sampson 2008; Xu et al. 2009). lurosaurian teeth are their small sizes and the absence of den- Morphotype 1 teeth, unlike those of neoceratosaurs, lack ticles. Similar teeth are known from several European and ornamentation in the tooth enamel. Ornamentation in the form North American sites. Teeth with similar characteristics from of wrinkles in the enamel is also commonly found in thero- the Hateg Basin were classified as indeterminate theropods pods like basal tetanurans, carcharodontosaurids and some (Antunes and Sigogneau-Russell 1991; Pol et al. 1992; Csiki tyrannosaurids (Brusatte et al. 2007). With the exception of and Grigorescu 1998; Garcia et al. 2000; López-Martínez et two teeth that are assigned to Daspletosaurus and Dilopho- al. 2001; Grigorescu 2003). saurus, Morphotype 1 teeth group in their own category in the discriminant analysis at generic/species level (SOM: Table Discussion.—Coelurosauria is a large clade that includes 2). At family level, only one of the teeth grouped with the many non-avian and avian theropod families. Most Creta- Neoceratosauridae, whereas most of the other Morphotype 1 ceous forms are small (under 2 m long), although some (some teeth showed stronger affinities to Tyrannosauridae. The sta- ornithomimids, most tyrannosauroids, and several dromaeo- tistically uncertain affinities of Morphotype 1 teeth suggests saurids) reached large sizes (Turner et al. 2007; Zanno and that they should be identified only as indeterminate theropods Makovicky 2013). Amongst small theropods, there are gen- until diagnostic postcranial material is found associated with era with distally curved teeth that do not have denticles and the teeth. are similar to those from Blasi 2B. These genera include the ornithomimosaur Pelecanimimus (Pérez-Moreno et al. Morphotype 2.—Teeth are shorter than those classified as Morphotype 1 and have more strongly curved crowns. How- 1994), troodontids like Byronosaurus and Mei (Norell et al. ever, measurements of the denticles are similar to those of 2000; Xu and Norell 2004), alvarezsaurids like Mononykus Morphotype 1, and these teeth could belong to the same and Shuvuuia (Chiappe et al. 1998; Suzuki et al. 2002), and taxon. Heterodonty in large theropods is widespread enough Mesozoic birds with teeth (Chiappe et al. 2002). Differentia- to encompass these two morphotypes in the same taxon tion can be difficult, although in birds there is a constriction (Smith 2005; Reichel 2010). It is also possible that Morpho- at the base of the crown (Sankey et al. 2002) that it has not type 2 could represent the juveniles of Morphotype 1. The been observed in any of the teeth in this study. curvatures of Morphotype 2 crowns are similar to those of the Statistically, these teeth grouped together in all three dis- juveniles of Tyrannosaurus rex described by Baszio (1997: criminant analyses (SOM: Table 2), although some speci- pl. 7: 94, 96, 101), whereas adult Tyrannosaurus rex teeth are mens grouped with Paronychodon because of similarities more similar to Morphotype 1. in size. However, they lack the characteristic longitudinal Statistically, Morphotype 2 groups on its own in the dis- ridges of Paronychodon. criminant analyses at species level (SOM: Table 2). At family level, the analysis reassigned the three teeth to Neovenatori- Family indet. dae (Benson et al. 2010). This clade is formed by some prob- Genus Paronychodon Cope, 1876 lematic allosauroids that survived into the Cretaceous. The Type species: Paronychodon lacustris Cope, 1876; Montana, USA; appearance of this clade in the Late Cretaceous of Europe Campanian. would have interesting palaeobiogeographic implications because the only known Late Cretaceous neovenatorid is ?Paronychodon sp. from South America. However, until more diagnostic ma- Fig. 3Q. 616 ACTA PALAEONTOLOGICA POLONICA 60 (3), 2015

ABC D

EF G H

I JK L

MNO P Q

Fig. 3. Theropod dinosaurs teeth from upper Campanian–Upper Maastrichtian, Spain. A. ?Pyroraptor olympius Allain and Taquet, 2000, DPM-MON-T1, Montrebei. B, C. ?Dromaeosauridae indet. B. MPZ2004/6, Blasi 2B. C. DPM-FON6-T2, Fontllonga 6. D–L. ?Richardoestesia sp., Laño. D. MCNA 14610. E. MCNA 14607. F. MCNA 14606. G. MCNA 14608. H. MCNA 14609. I. MCNA 14611. J. MCNA 14568. K. MCNA 14607. L. MCNA 14619. M–P. Coelurosauria indet. M. DPM-MON-T6, Montrebei. N. DPM-MON-T3, Montrebei. O. MPZ98/80, Montrebei. P. MPZ98/82, Blasi 2B. Q. ?Paron- ychodon sp., MPZ98/76, Blasi 2B. All lateral views. Scale bars 1 mm.

1991 Euronychodon portucalensis Antunes and Sigogneau-Russell, Material.—Three teeth (MPZ98/76–78) from Blasi 2B (Tremp 1991: 118–119, fig. 15A. Formation, late Maastrichtian) (Fig. 3Q; SOM: Table 1). 2001 cf. Euronychodon sp.; López-Martínez et al. 2001: 49, 51, fig. 9B, C, D. Description.—Paronychodon teeth are relatively tall, elon- 2004 cf. Euronychodon sp.; Torices et al. 2004: 72–73. gate and curve gently (Currie et al. 1990). Usually, the pos- TORICES ET AL.—LATE CRETACEOUS THEROPOD DINOSAURS FROM SPAIN 617

3

2

1

-2.5 -2 -1.5 -1 0.5 1 1.5

PC 2 -0.5

-1

-2

Dromaeosaurus PC 1 ?Richardoestesia sp.

Saurornitholestes Colurosauria indet.

Pyroraptor olympius Richardoestesia Troodon

Theropoda indet. Morphotype 2 ?Pyroraptor olympius ?Dromaeosauridae indet.

Theropoda indet. Morphotype 1 Paronychodon ?Paronychodon sp.

Fig. 4. Principal component analysis of the South Pyrenees Basin sample and the Royal Tyrrell Museum of Palaeontology sample; a chart displaying two first principal components, PC1 and PC2. terior carina is more or less straight and the anterior one is such as crocodyliforms. Rauhut and Zinke (1995) suggested convex in lateral view, but both carinae can be convex. The that Paronychodon teeth can be attributed to Pelecanimi- enamel is sometimes folded, and three or more longitudinal mus, but this seems unlikely (Rauhut 2002). Rauhut (2002) ridges appear on one or both flanks of the teeth. Denticles are proposed that they could belong to birds. The longitudinal never present (Currie et al. 1990). ridges in Paronychodon teeth have been attributed to growth Three teeth from Blasi 2B are referred to ?Paronycho- abnormalities (Currie et al. 1990). Other authors consid- don sp. They have similar measurements and morphological er them normal and refer “paronychodontids” to primitive characteristics to the ones described as Coelurosauria indet. maniraptoriform theropods, as close relatives of troodontids, They are small, and their heights are less than 3 mm, their ornithomimosaurs, or even birds (Zinke and Rauhut 1994; FABL varies between 1.21 and 1.49, and their basal widths Csiki and Grigorescu 1998; Rauhut 2002). range between 0.60 and 0.84. They lack denticles on their Teeth from our sample are similar to Paronychodon lacustris anterior and posterior carinae, but longitudinal ridges can from the Upper Cretaceous of North America that have been be observed on the labial and lingual sides of their crowns. classified as Paronychodon lacustris (Currie et al. 1990) and Discussion.—Paronychodon is an enigmatic theropod taxon group together perfectly (Fig. 4). They are also similar to a that has teeth ornamented with longitudinal ridges (Canudo Portuguese tooth classified as Euronychodon portucalensis and Ruiz-Omeñaca 2003). The position of this genus within (Antunes and Sigogneau-Russell 1991). Other similar teeth Theropoda is uncertain. There is no clear reason to assign have been identified as cf. Paronychodon or cf. Euronycho- it to Dromaeosauridae, as Antunes and Sigogneau-Russell don in French, Romanian, and Spanish sites. (1991) tentatively suggested. On the other hand, the assign- North American teeth of Paronychodon are indistinguish- ment of Paronychodon (as a nomen dubium) to Troodontidae able from Euronychodon, so it is probable that Euronycho- by Osmólska and Barsbold (1990) and Makovicky and Norell don is a synonym (Rauhut 2002). Pyrenean teeth previously (2004), is probably based on the slight constriction between classified as cf. Euronychodon (Torices et al. 2004) are there- crown and root; this is insufficient because this character fore renamed here as ?Paronychodon. can be found in other theropods (Zinke and Rauhut 1994), in Statistically, these teeth grouped consistently in all three toothed birds (Currie 1987), and in some other vertebrates, discriminant analyses. Although one specimen grouped with 618 ACTA PALAEONTOLOGICA POLONICA 60 (3), 2015 indeterminate coelurosaurians because of similar size, its as- mm. The morphology of each anterior denticle is also rectan- signment is certain because of the presence of longitudinal gular in lateral view, but the denticles are only found on the ridges in its enamel (SOM: Table 2). apical half of the carina. The material from Vicari 4 consists of one entire tooth, Family indet. and the apex of another. The complete tooth has a convex Genus Richardoestesia Currie, Rigby, and Sloan, 1990 anterior carina and a concave posterior one in lateral view. Both teeth are laterally compressed with basal widths that Type species: Richardoestesia gilmorei Currie, Rigby, and Sloan, 1990; Dinosaur Provincial Park; Campanian. are about half of the FABLs. Posterior denticle density is 9.5 denticles/mm. The morphology of each denticle is rectan- ?Richardoestesia sp. gular, and some of them (including DPM-VIR4-T7) do not Fig. 3D–L. show any wear. The 56 teeth of Laño constitute most of the Richardoeste- 2003 Dromaeosauridae indet. 4; Torices 2002: 141. 2004 cf. Dromaeosauridae morphotype 8; Torices et al. 2004: 73. sia sample. They consist of complete teeth and fragments 2004 cf. Dromaeosauridae indet. 4; Torices et al. 2004: 73. that retain only the middle parts of the teeth, the apices, or 2005 Maniraptoriformes indet.; Canudo et al. 2005: 34. the posterior carinae. They all have denticles on the posterior carinae, with densities between 6.5 and 11.4 denticles/mm. In Material.—Four teeth (MPZ98/72–74, 2004/7) from Blasi sixteen of the specimens, anterior denticles are also present, 2B (Tremp Formation, late Maastrichtian); sixty-four isolat- with densities between 8.1 to 16.3 denticles/mm. The shape ed teeth (MCNA 14566–14621) from Laño (lateral equiva- of each posterior denticle is rectangular, and in some cases a lent of Sedano Formation, upper Campanian–lower Maas- slight tilt toward the apex of the tooth can be observed. The trichtian); two teeth (DPM-MON-T5, T9) from Montrebei morphology of each anterior denticle is also rectangular, but (Tremp Formation, upper Campanian–lower Maastrichtian); its overall size is much smaller than an equivalent poste- two teeth (DPM-VIR4-T6, T7) from Vicari 4 (Tremp Forma- rior denticle. The morphologies of the tooth crowns cover tion, upper Campanian) (Fig. 3D–L; SOM: Table 1). the three types: (i) biconvex teeth; (ii) teeth with convex Description.—Richardoestesia specimens are small thero- anterior carinae and straight posterior ones; and (iii) teeth pod teeth (crown height between 1.4 and 5.1 mm, FABL with convex anterior carinae and concave posterior ones. between 1 and 5.1 mm, and basal width between 0.4 and 2.2 Those with strongly concave posterior borders are usually mm) with posterior denticle densities between 6 and 11.4 lower in height than those with straight edges, which are denticles/mm. Sixteen of the sixty four specimens have an- more elongate vertically. The lower heights suggest that this terior denticles that are smaller than the posterior ones, and morphology may represent posterior maxillary or posterior with densities between 8.1 and 16.3 anterior denticles/mm. dentary teeth. Denticles are small and rectangular. In some of the teeth, it is Teeth identified as Richardoestesia in the Royal Tyrrell possible to see that the posterior denticles are tilted slightly Museum of Palaeontology (Drumheller, Canada) collections towards the apices of the teeth, which is characteristic of the were measured. The height varies from 2.5 to 10.8 mm, the genus Richardoestesia. FABL varies between 1.37 and 4.67 and basal width ranges The morphologies of the teeth have great variability, from from 0.66 to 2.83 mm. Denticle densities vary from 5 to 12.5 teeth with convex anterior borders and concave posterior posterior denticles/mm, and 5 to 10 anterior denticles/mm borders to teeth with convex anterior borders and straight when anterior denticles are present. Each of these measure- posterior borders. In some teeth, both edges are slightly con- ments overlap perfectly with those of the Pyrenean sample vex, and the teeth resemble isosceles triangles. (Figs. 4 and 5). The four specimens of Richardoestesia from the Blasi Discussion.—The morphologies and measurements of many 2B site have almost straight (slightly biconvex) carinae, and Spanish specimens fall within the ranges of variability of compressed basal sections (basal widths are about half their teeth from North America that are identified as Richardoeste- FABLs). The tips of three of the teeth are broken and one of sia (Currie et al. 1990) (Figs. 4 and 5). Richardoestesia gil- them is slightly worn, and their roots are not preserved. Their morei was described by Currie et al. (1990) on the basis of a denticles are rectangular. pair of lower jaws containing a replacement tooth. Isolated The two specimens from Montrebei have convex anterior teeth were assigned to this species based on the character- borders and straight posterior ones. They are compressed istics of the denticles, particularly their small size. Richar- lateromedially and their basal widths are nearly half of the doestesia denticles are the smallest seen in any of the small corresponding FABLs. The tips show wear that in DPM- theropods from the Upper Cretaceous. Usually they do not MON-T5 affects the anterior carina, and there is some break- have anterior denticles, but when present, they are even tinier age affecting the posterior carina in both teeth. The density than the posterior denticles. of posterior denticles ranges from 7.8 to 9 denticles/mm. In Richardoestesia collections from the Late Cretaceous The morphology of each denticle is rectangular. Only DPM- of Canada and the USA, various researchers (Currie et al. MON-T9 has denticles on the anterior carina that are smaller 1990; Baszio 1997; Sankey et al. 2002) have observed two than the posterior ones; the density is 11.5 anterior denticles/ different morphologies: (i) tall and straight teeth, and (ii) TORICES ET AL.—LATE CRETACEOUS THEROPOD DINOSAURS FROM SPAIN 619

Dromaeosaurus 12 Saurornitholestes Troodon Richardoestesia 10 ?Richardoestesia sp. ?Dromaeosauridae indet. ?Pyroraptor olympius 8 Pyroraptor olimpus “Richardoestesia-like”

6

4

Posterior denticles / mm

2

0

0 5 10Height 15 20 25 Fig. 5. Bivariate analysis comparing height (in mm) against posterior denticles per millimeter, ?Dromaeosauridae, ?Pyroraptor olympius, and ?Richar- doestesia from the South Pyrenees area are compared against a sample of Dromaeosaurus, Saurornitholestes, Richardoestesia, and Troodon from the collections of the Royal Tyrrell Museum of Palaeontology, the Richardoestesia-like tooth from the site of Suterranya, Catalonia, Spain (Prieto-Márquez et al. 2000) and Pyroraptor olympius Allain and Taquet, 2000 from Provence (Ronan Allain, personal communication 2013). shorter and curved teeth. Currie et al. (1990) considered the Cretaceous of Romania (Codrea et al. 2002; Weishampel et al. possibility that these two morphologies represent positional 2010). The measurements and morphological characteristics variation along the dental series, with the straighter ones of the teeth and their denticles are also perfectly compatible coming from the anterior parts of the jaws. However, they with those in the Spanish sample from the Upper Cretaceous. also alluded to the possibility that the two morphotypes rep- A tooth from L’Abella in the Upper Cretaceous of the resented two different taxa. Baszio (1997) preferred the latter Pyrenees was referred to as Richardoestesia-like (Prieto- option, and he distinguished the straight teeth as a distinct Márquez et al. 2000). This tooth has a FABL of 3.1 mm and species (Richardoestesia sp.) that Sankey (2001) named a posterior denticle density of 7 denticles/mm. The anterior Richardoestesia isosceles. The latter paper also recogniz- edge is broken, so it is impossible to know whether or not it es differences in the morphologies of the denticles between possessed anterior denticles. The denticles are rectangular. Richardoestesia gilmorei and Richardoestesia isosceles. The It is a taller tooth (15.3 mm) than the rest of the specimens denticles of Richardoestesia isosceles are more square and assigned to the genus (the average height is 4.9 mm) (Fig. 5). lack interdenticular spaces (Sankey et al. 2002). Although In the discriminant analysis performed, this tooth is clearly Richardoestesia gilmorei and Richardoestesia isosceles form isolated by its height from both the North American and two distinct morphological groups, they are not distinguish- Pyrenean samples (SOM: Table 2), which suggests it might able from each other when considering variables such as be a new taxon. crown height and FABL. In principle, the differences in the general morphology of the tooth that Baszio (1997) used to Dromaeosauridae Matthew and Brown, 1922 distinguish Richardoestesia sp. teeth from those of Richar- Genus Dromaeosaurus Matthew and Brown, 1922 doestesia gilmorei are not sufficient proof because morpho- logical variability within even a single theropod jaw can Type species: Dromaeosaurus albertensis Matthew and Brown, 1922; be remarkable (Smith 2005, Reichel 2010). Company et al. Red Deer River, Campanian. (2005) proposed that Richardoestesia isosceles might be a ?Dromaeosauridae indet. ziphodont crocodyliform. 2002 Dromaeosauridae indeterminate 2; Torices 2002: 141. Teeth identified as cf. Richardoestesia have been recov- 2004 cf. Dromaeosauridae morphotypes 2, 5 and 6; Torices et al. 2004: ered from the Upper Jurassic of Portugal, the Lower Cre- 73. taceous of Spain (Zinke 1998; Rauhut 2002) and the Upper 2005 Maniraptoriforme indet.; Canudo et al. 2005: 34. 620 ACTA PALAEONTOLOGICA POLONICA 60 (3), 2015

Material.—Four specimens: MPZ2004/6 from Blasi (Arén of the tooth, and then twists onto the lingual surface (Currie Formation, upper Maastrichtian); MCNA 14622 from Laño et al. 1990). The absense of this feature in the Spanish teeth (lateral equivalent of Sedano Formation, upper Campanian– suggests that they cannot be assigned to the genus Dromaeo- lower Maastrichtian); DPM-FON6-T2 from Fontllonga 6 saurus. They are designated here as indeterminate dromeo- (Tremp Formation, lower Maastrichtian); DPM-FIG2-T1+T2 saurid teeth. from Figuerola 2 (Tremp Formation, upper Campanian) (Figs. 3B, C, SOM: Table 1). Genus Pyroraptor Allain and Taquet, 2000 Description.—The measurements of the teeth of indetermi- Type species: Pyroraptor olympius Allain and Taquet, 2000, Provence. nate dromaeosaurids vary from a minimum crown height of ?Pyroraptor olympius Allain and Taquet, 2000 3.4 mm (as estimate from a broken specimen) to 17.5 mm, 2002 Dromaeosauridae indeterminate 3; Torices 2002: 141–143. FABLs of 2.6 to 16 mm, and basal widths between 1.3 and 2004 cf. Dromaeosauridae indeterminate Morphotype 3; Torices et al. 5.8 mm. Crown shapes in lateral or medial views corre- 2004: 73. spond to two types: (i) biconvex borders or (ii) a convex Material.—Four teeth (MCNA 14623–14626) from the Laño anterior border and a straight posterior one. These laterally (lateral equivalent of Sedano Formation, upper Campanian– compressed teeth have denticulate carinae. Denticle densities lower Maastrichtian); one (DPM-MON-T1) from Montrebei vary between 3.5 to 8.1 anterior denticles/mm and 2.5 to (Tremp Formation, upper Campanian–lower Maastrichtian) 4.9 mm posterior denticles/mm. The denticles are rectangu- (Figs. 2B, C, 3A) lar, slightly rounded and chisel-like (Currie et al. 1990). The indeterminate dromaeosaurid from Laño consists of Description.—The teeth assigned to ?Pyroraptor olympius only the tip of a tooth that has a convex anterior border and are laterally compressed with convex anterior borders and straight posterior one. Denticle densities are 8.1 anterior den- concave posterior ones. Their heights vary between 3.8 and ticles/mm and 4.9 posterior denticles/mm. 8 mm, FABLs between 2.5 and 5.4 mm, and basal widths The specimen from Fontllonga 6 is biconvex in mor- range between 1.4 and 2.4 mm. Anterior denticle densities phology and has large denticles with a density of 3 posterior vary between 8 and 9.8 denticles/mm and posterior denticle denticles/mm. The tooth from Figuerola 2 lacks enamel, al- densities are 6 to 6.5 denticles/mm. The denticles are square. though it is possible to see traces of the denticles and measure Three of the teeth were found in situ within a dentary bone their density (2.5 posterior denticles/mm). fragment and the other one was found isolated but in asso- The teeth from Blasi have convex anterior borders and ciation with the dentary fragment (MCNA 14623–14626). straight posterior ones. Their anterior denticle densities vary In this fragment, it can be seen that the interdental plates between 3.5 and 5 denticles per millimeter, whereas their pos- are fused together, which is characteristic of dromeosaurids terior denticle densities vary between 2.9 and 3 denticles/mm. and has been observed in Deinonychus, Dromaeosaurus, and Saurornitholestes (Currie et al. 1990; Currie 1995). This Discussion.—All Dromaeosaurus teeth in the collections of character has also been observed in some larger theropods the Royal Tyrrell Museum of Palaeontology and those from like carcharodontosaurids, megalosaurids and neovenatorids the holotype (AMNH 5356) of Dromaeosaurus albertensis (Brusatte et al. 2008, 2012; Benson 2010). Characteristics of Matthew and Brown, 1922 were measured. The height of the these teeth are similar to those of the dromaeosaur Pyroraptor teeth of this sample varies between 5.33 and 26 mm, FABL olympius (Allain and Taquet 2002) in the density of denticles varies between 2.17 and 8.83, and basal width ranges from (6 posterior denticles/mm) and the square denticle morphol- 1.67 to 9.17. Anterior denticle density is 1 to 8.75 denticles/ ogy, so the probability is high that our sample could belong mm, whereas posterior denticle density is 1 to 6.25 denticles/ to the same taxon. mm. The measurements of the teeth from the Pyrenean sam- ple fall within the ranges of these measurements and those Discussion.—Statistically, the teeth from the Pyrenean sam- described in the literature (Currie et al. 1990; Baszio 1997; ple grouped together with Pyroraptor teeth in the three dis- Sankey et al. 2002). The rectangular, chisel-like morpholo- criminant analyses performed (SOM: Table 2). gies of the denticles match those described for dromaeosau- Therefore, considering the characters that identify our rids (Figs. 4, 5). sample as a dromaeosaurid, the similarity of morphological The discriminant analyses suggest that all teeth fall in the and numerical data to those of Pyroraptor teeth and the cor- Dromaeosauridae; two of them are grouped with Dromaeo- rect assignation by the discriminant analyses of our sample saurus teeth and one is placed with ?Pyroraptor teeth (SOM: to Pyroraptor, these teeth are best identified as ?Pyroraptor Table 2). The last tooth was not identified as Pyroraptor olympius. because its denticles resemble the taller, narrower ones of dromaeosaurids than the lower, broader ones of Pyroraptor. It should be noted, however, that there is a diagnostic char- Results acteristic of Dromaeosaurus that is not found in any of the Spanish teeth. The anterior carina of each tooth of Dromaeo- Principal components analysis of the log-transformed data of saurus extends along the midline from the apex to mid-height the Pyrenean and North American samples indicates that two TORICES ET AL.—LATE CRETACEOUS THEROPOD DINOSAURS FROM SPAIN 621 components explain the variance of the data (Fig. 4). PC1 denticles/mm) of Dromaeosaurus, but the morphologies shows a heavy loading in the tooth size variables (height, of their respective denticles are different (?Dromaeosauri- FABL, basal width) and PC2 shows a heavier loading in dae indet. are taller and narrower than those of ?Pyroraptor the denticle variables. Graphically the representation of the olympius). For this reason it is reasonable to assign these two components groups the different taxa with some overlap morphotypes to two different taxa. Teeth identified as ?Rich- especially between Dromaeosaurus, Saurornitholestes, and ardoestesia are distinctive in size and denticle density. They Troodon. The rest of the taxa are grouped better allowing group well statistically in the discriminant analysis with the comparison of the two samples and helping the identification North American sample of Richardoestesia. Teeth described of the Pyrenean sample. as ?Paronychodon teeth have diagnostic longitudinal ridges Bivariate analyses show that there is a strong correla- that separate them from smaller, non-denticulate teeth. Apart tion (significant at 0.01) between the five variables—crown from the teeth studied in this work, another tooth needs to height, fore-aft-basal length (FABL), basal width, posteri- be considered. It was found at the Suterranya site, near the or denticle density, and anterior denticle density. In Fig. 5, Vicari 4 site, described by Prieto-Márquez et al. (2000) and the relationship between tooth size, represented by crown determined to be a Richardoestesia-like theropod. This tooth height, and denticle size, represented by number of posterior was included in the discriminant analysis and did not group denticles per millimeter, is shown for small theropods from with any of the taxa described in this paper because of its size the two samples. The different taxa are grouped with some (Fig. 5; SOM: Table 2), suggesting it may belong to another overlap, especially between Dromaeosaurus, Saurornitho- theropod taxon. The remains of a possible ornithomimosaur lestes, and Troodon. In this case, the different morphology were identified from Laño (Astibia et al. 1990), and this may of their denticles is the key for their identification in spite of be evidence of another theropod taxon in the area that may the overlapping of the numerical data. not be represented by teeth. The discriminant analyses gave poor results, correctly A total of eight taxa of theropods are present in the South identifying only 58.5% of specimens at species level and Pyrenees Basin. There are six taxa present at Laño, and 67.4% at family level. The reason for these poor results is eight in the central Pyrenees sites (four taxa in Blasi, one in the enormous amount of overlap between the teeth of large Figuerola 2, one in Fontllonga 6, three in Montrebei and three theropods (especially between tyrannosaurids). There is inVicari 4) (SOM: Table 3). some overlap in the small theropods but to a lesser degree. The change in diversity of theropod dinosaurs through The third discriminant analysis was performed for the Pyre- the Late Cretaceous (late Campanian–late Maastrichtian) can nean sample, and this gave better results, with the percentage be analysed from the sites of South Pyrenees Basin (SOM: of correctly identified species as 79.2% (SOM: Table 2). Table 3). Figuerola 2, Suterranya, and Vicari 4 represent up- per Campanian deposits, where four theropod taxa are pres- ent. Laño and Montrebei represent upper Campanian–lower Discussion Maastrichtian deposits, where there are six taxa represented. In the lower Maastrichtian, there is only one site, Fontllonga Many newly discovered teeth from the Campanian–Maas- 6, where only one kind of theropod is present. And finally, in trichtian of the South Pyrenees Basin significantly increase the upper Maastrichtian (represented by deposits from Blasi our knowledge about theropod diversity in the Iberian Pen- 1, 2B, and 3), there are five taxa of theropods. insula at the end of the Cretaceous. Comparing the diversity of Campanian–Maastrichtian In this work, seven morphotypes of theropod teeth are theropods in the Iberian Peninsula with the rest of Europe identified from the South Pyrenees Basin. Two morphotypes at the same time, the faunal associations follow similar pat- of Theropoda indet. are separated in the discriminant anal- terns. Based on teeth, a large theropod and a series of small yses even though their denticle densities and denticle mor- theropods (typically dromaeosaurids and other theropods phologies are nearly identical. The main difference between with teeth that lack denticles) can be recognized. Compared them is crown shape. Heterodonty has been observed in the to the rich associations of France, Portugal and Romania jaws of large theropods (Smith 2005; Reichel 2010). Elon- (teeth and other fossils), the Spanish fauna of theropods is gate, more rounded teeth characterize the anterior parts of the as good or better in terms of number of taxa represented. jaws, and shorter, more recurved teeth belong to the posterior France has six taxa of theropods, Portugal has five, Hunga- parts (Smith 2005). For this reason, the most conservative ry has one taxon in the Early Santonian, and Romania has approach would be to assign these two morphotypes to a nine (Buffetaut et al. 1986; Antunes and Sigogneau-Russell single taxon. Two morphotypes of dromeosaurid teeth have 1991; Buffetaut and Le Loeuff 1991, 1997; Le Loeuff and been identified as ?Dromaeosauridae indet. and ?Pyrorap- Buffetaut 1991, 1998; Csiki and Grigorescu 1998; Pere- tor olympius. The two morphotypes group separately in the da-Suberbiola 1999b; Allain and Taquet 2000; Garcia et al. discriminant analysis and ?Pyroraptor teeth group togeth- 2000; Allain and Pereda-Suberbiola 2003; Bonde and Chris- er with the teeth from the holotype of Pyroraptor giving tiansen 2003; Grigorescu 2003; Jagt et al. 2003; Csiki et al. strength to this assignment. The variabilities of their denticle 2010; Weishampel et al. 2010; Brusatte et al. 2013) (SOM: densities both fall within the range of variability (2.5–6.5 Table 4). 622 ACTA PALAEONTOLOGICA POLONICA 60 (3), 2015

Most of the studies on theropod tooth assemblages in Eu- Maastrichtian taxa (Theropoda indet., Coelurosauria indet., rope have been done using qualitative (but not quantitative) ?Richardoestesia sp., ?Dromaeosauridae indet., ?Pyroraptor comparisons, mostly with North American faunas. Only Ösi olympius, and ?Ornithomimosauria indet. described by Astib- et al. (2010) used statistical analyses to help identify Hun- ia et al. 1990). Although only one taxon (?Dromaeosauridae garian theropods (Santonian). In terms of composition, there indet.) appears in the early Maastrichtian, this is a sampling are many similarities between different European sites, based problem that cannot be resolved at this time. The presence on the presence of dromaeosaurids, ?Paronychodon, other of five theropod taxa (indeterminate theropod, indeterminate coelurosaurians, and a large theropod. However, they differ coelurosaurian, ?Paronychodon sp., ?Richardoestesia sp., from the South Pyrenees Basin in that most of these other ?Dromaeosauridae indet.) in the late Maastrichtian suggests deposits seem to lack identifiable Richardoestesia teeth. In there was no significant decrease in the theropod diversity the Hateg basin of Romania, Codrea et al. (2002) mention at the end of the Cretaceous in this region (SOM: Table 3). three teeth similar in morphotype to Richardoestesia, but ul- These results agree with Lillegraven and Eberle (1999), who timately classified them as Theropoda incertae sedis. In the observed that diversity remained high in North America until Spanish deposits, specimens of Troodon or troodontids have the uppermost levels, although they do not rule out a gradual not been recognized, but they have been described from the (non-catastrophic) extinction of the dinosaurs. Sheehan et al. Portuguese and Romanian deposits (SOM: Table 4). (2000) and Lyson et al. (2011) found similar results in the up- In the South Pyrenees Basin and most other European permost three meters of the Hell Creek formation. However, outcrops, the remains (mostly teeth, but some postcranial the diversity of dinosaurs in North America is considerably material) of large theropods, have been found (Buffetaut et less in the Hell Creek Formation than it is in the Judith River al. 1986; Antunes and Sigogneau-Russell 1991; Buffetaut Formation (Weishampel et al. 2004). and Le Loeuff 1991, 1997; Le Loeuff and Buffetaut 1991, 1998; Pereda-Suberbiola 1999b; Allain and Taquet 2000; Garcia et al. 2000; Allain and Pereda-Suberbiola 2003; Conclusions Bonde and Christiansen 2003; Jagt et al. 2003). It is worth mentioning that no large theropod teeth have been recovered The study of 142 isolated teeth from the Campanian–Maas- in the Hateg Basin of Romania. Nopcsa (1902) described trichtian of the South-Pyrenean Basin suggests six species some theropod teeth as Megalosaurus hungaricus but these of toothed theropods (five small, one large) were present in teeth came from Borod Basin (possibly Santonian) and not the region. The taxa identified include two morphotypes of from the Hateg Basin. Unfortunately, the specimens have an indeterminate theropod (which could correspond to juve- been lost (Csiki and Grigorescu 1998; Csiki et al. 2010; Bru- niles and adults of the same species, or tooth row variation), satte et al. 2013). The absence of large theropods from the Coelurosauria indet., ?Paronychodon sp., ?Richardoestesia Hateg fauna has been attributed to the palaeoenvironments sp., ?Dromaeosauridae indet., ?Pyroraptor olympius. Other during the Cretaceous (Csiki and Grigorescu 1998; Csiki et two taxa, a Richardoestesia-like form (Prieto-Márquez et al. al. 2010; Brusatte et al. 2013). 2000) and a possible ornithomimosaur are added to the final The establishment of a chronostratigraphical framework diversity. In total, there are eight theropod taxa present in the (from Upper Campanian to Upper Maastrichtian) for thero- South Pyrenees Basin at the end of the Cretaceous. pod teeth from the Pyrenees Basin provides another tool for At least five families appear to be present. The indetermi- evaluating the ages of palaeontological sites. Palaeoenviron- nate theropod teeth identified as Morphotype 1 and Morpho- mentally, the Arén and Tremp Formations were formed by type 2 represent a large theropod of uncertain affinities. Sta- a barrier island and coastal lagoon system. The taphonomic tistically these teeth show more affinities at the family level in characteristics are roughly equivalent although each of the the analyses with tyrannosaurids, and only one tooth grouped sites in these formations has its own peculiarities. The sites with Neoceratosauridae. A possible ornithomimosaurid is were prospected and sampled with approximately the same represented by some phalanges. The family Dromaeosauri- intensity, although the Fontllonga 6 sample is inadequate dae is represented by teeth identified as ?Dromaeosauridae because of its small size. The Laño deposits were formed in indet. and ?Pyroraptor olympius. Specimens identified as an alluvial system and the palaeoenvironmental taphonomic ?Richardoestesia sp. represent Maniraptoriformes, but the characteristics are different from other Pyrenean sites. The family of this genus is still uncertain. Similarly, the families intensity of sampling in the Laño deposits was higher and represented by the indeterminate coelurosaurian and ?Paron- produced a larger sample. Even so, most of the taxa that ap- ychodon sp. are unknown. pear in Laño are present in the rest of Pyrenean sites and fit With this study the number of theropod taxa known from within the framework of changes over time. the South Pyrenees Basin is exponentially increased. This In the South Pyrenees Basin, it appears that overall di- shows the value of isolated teeth to reconstruct the compo- versity increases from four theropods (an indeterminate sition of dinosaur palaeofaunas when other more complete coelurosaurian, ?Richardoestesia sp., a Richardoestesia-like material is not present allowing us to make interpretations form described by Prieto-Márquez et al. 2000, and an in- about the evolution of their diversity through time. In this determinate dromeosaurid) in the late Campanian to six case, apparently theropod diversity in the north of Spain TORICES ET AL.—LATE CRETACEOUS THEROPOD DINOSAURS FROM SPAIN 623 does not experience a significant decline at the end of the Astibia, H., Buffetaut, E., Buscalioni, A., Cappetta, H., Corral, J.C., Estes, Cretaceous (Campanian and Maastrichtian). R., García-Garmilla, F., Jaeger, J.J., Jimenez Fuentes, E., Le Loeuff, J., Mazin, J., Orue-Etxebarria, X., Pereda-Suberbiola, X., Powell, J.E., Rodríguez-Lázaro, J., Sanz, J.L., and Tong, H. 1990. The fossil verte- brates from Laño (Basque Country, Spain); new evidence on the compo- sition and affinities of the Late Cretaceous continental faunas of Europe. Acknowledgements Terra Nova 2: 460–466. The authors want to thank Nieves López-Martínez, who initiated and Astibia, H., Garcia-Garmilla, F, Orue-Extebarria, X., Rodríguez-Lazaro, J, encouraged this study, and was its driving force up until her premature Buscalioni, A.D., Sanz, J.L., and Jimenez-Fuentes, E. 1987. The Cre- taceous–Tertiary boundary in a sector of the south limb of the Miran- death; this paper is dedicated to her memory. We acknowledge Luis da-Trevino Synclinal: The first appearance of Chelonia and Archosau- Ardèvol (Geoplay, Tremp, Spain), Otto Kälin and Margarita Díaz-Mo- ria in the Basque Country. Cretaceous Research 8: 15–27. lina (both DPM) for their help in the field, José Ignacio Ruiz-Omeñaca Baceta, J.I., Pujalte, V., and Orue-Etxebarria, X. 1999. The vertebrate fossil - (Museo del Jurásico de Asturias, Colunga, Spain) for early discussions, bearing sites of the Laño quarry (Basque-Cantabrian Region): stratigra- Don Brinkman and Brandon Strilisky (both Royal Tyrrell Museum of phical and paleogeographical context. Estudios del Museo de Ciencias Palaeontology, Drumheller, Canada) for allowing access to the Roy- Naturales de Álava 14, Número Especial 1: 13–28. al Tyrrell Museum collections, and Ronan Allain (Muséum National Baszio, S. 1997. Systematic Palaeontology of Isolated Dinosaur Teeth d’Histoire Naturelle, Paris, France) for access to Pyroraptor olympius from the Latest Cretaceous of South Alberta, Canada. Courier For- data. We thank Victoria Arbour (North Carolina Museum of Natural schungsinstitut Senckenberg 196: 33–77. Sciences, Raleigh, USA) for English revision. We thank Steve Brusatte Benson, R.B.J. 2010. Description of Megalosaurus bucklandii (Dinosau- (University of Edinburgh, UK) and Nate Smith (Howard University, ria: Theropoda) from the Bathonian of the United Kingdom and the Washington, USA) for their helpful comments that greatly improved relationships of Middle Jurassic theropods. Zoological Journal of the this paper. We thank Michael Benton (University of Bristol, UK) for Linnean Society 158: 882–935. his help with comments and editorial work. Funds were provided by Benson, R.B.J., Carrano M.T., and Brusatte, S.L. 2010. A new clade of projects CGL 2006-04646, CGL2010-16447, and CGL 2009-09000 archaic large-bodied predatory dinosaurs (Theropoda: Allosauroidea) (Spanish Ministry of Education and Science and the Ministry of Sci- that survived to the latest Mesozoic. Naturwissenschaften 97: 71–78. ence and Innovation), Research Group no. 910161, Registro geológico Blain, H.A., Canudo, J.I., Cuenca-Bescós, G., and López-Martínez, N. de períodos críticos: factores paleoclimáticos y paleoambientales (Uni- 2010. Amphibians and squamate reptiles from the latest Maastrichtian (Upper Cretaceous) of Blasi 2 (Huesca, Spain). Cretaceous Research versidad Complutense de Madrid, Autonomous Community of Madrid, 31 (4): 433–446. Spain), the European Regional Development Fund, the Government Bonaparte, J.F., Novas, F.E., and Coria, R.A. 1990. Carnotaurus sas- of Aragón (“Grupos Consolidados” and “Dirección General de Patri- trei Bonaparte, the horned, lightly built carnosaur from the middle monio”). Research of XPS supported by the Ministerio de Economía Cretaceous of Patagonia. Contributions in Science, Natural History y Competitividad (CGL2010-18851/BTE) and the Gobierno Vasco/EJ Museum of Los Angeles County 416: 1–42. (research groups IT-320-10 and IT834-13). AT is funded by FECYT Bonde, N. and Christiansen, P. 2003. New dinosaurs from Denmark. Comp- and the program “Ayudas para la movilidad postdoctoral en centros tes Rendus Palevol 2: 13–26. extrajeros”. Brusatte, S.L., Benson, R.B.J., and Hutt, S. 2008. The osteology of Neove- nator salerii (Dinosauria: Theropoda) from the Wealden Group (Bar- remian) of the Isle of Wight. Monograph of the Palaeontographical Society 162 (631): 1–166. References Brusatte, S.L., Benson, R.B.J., and Xu, X. 2012. A reassessment of Kel- mayisaurus petrolicus, a large theropod dinosaur from the Early Creta- Accarie, H., Beaudoin, B., Dejax, J., Friès, G., Michard, J.-G., and Taquet, ceous of China. Acta Palaeontologica Polonica 57: 65–72. P. 1995. Découverte d’un Dinosaure théropode nouveau (Genusaurus Brusatte, S.L., Benson, R.B.J., Carr, T.D., Williamson T.E., and Sereno sisteronis n. g., n. sp.) dans l’Albien marin de Sisteron (Alpes de Haute- P.C. 2007. The systematic utility of theropod enamel wrinkles. Journal Provence, France) et extension au Crétacé inférieur de la lignée cérato- of Vertebrate Paleontology. 27: 1052–1056. saurienne. Comptes rendus de l’Academie des Sciences, Paris, II 320: Brusatte, S.L., Vremir, M., Csiki-Sava, Z., Turner, A.H., Watanabe, A., Er- 327–334. ickson, G.M., and Norell, M.A. 2013. The osteology of Balaur bon- Allain, R. 1998. Un nouveau gisement de vertébrés continentaux du Crétacé doc, an island-dwelling dromaeosaurid (Dinosauria: Theropoda) from supérieur du bassin de l’Arc (Bouches-du-Rhône). Description systé- the Late Cretaceous of Romania. Bulletin of the American Museum of ma tique et implications paléobiogéographiques. 141 pp. Unpublished Natural History 374: 1–100. Mémoire de DEA, Université de Montpellier-2, Montpellier. Buffetaut, E. and Le Loeuff, J. 1991. Late Cretaceous dinosaur faunas of Allain, R. and Pereda-Suberbiola, X. 2003. Dinosaurs of France. Comptes Europe: some correlation problems. Cretaceous Research 12: 159–176. Rendus Palevol 2: 27–44. Buffetaut, E. and Le Loeuff, J. 1997. Late Cretaceous dinosaurs from the Allain, R. and Taquet, P. 2000. A new genus of Dromaeosauridae (Dino- foothills of the Pyrenees. Geology Today 13 (2): 60–68. sauria, Theropoda) from the Upper Cretaceous of France. Journal of Buffetaut, E., Marandat, B., and Sigé, B. 1986. Discovery of deinonycho- Vetebrate Paleontology 20: 404–407. saur Teeth (Saurischia, Theropoda) in the Upper Cretaceous of South- Allain, R., Tykoski, R., Aquesbi, N., Jalil, N.E., Monbaron, M., Russell, ern France. Comptes rendus de l’Académie des Sciences de Paris, II, D., and Taquet, P. 2007. A basal abelisauroid from the late Early Juras- 303: 1393–1396. sic of the High Atlas Mountains, Morocco, and the radiation of cerato- Buffetaut, E., Mechin, P., and Mechin-Salessy, A. 1988. Un dinosaure saurs. Journal of Vertebrate Paleontology 27: 610–624. théropode d’affinités gondwaniennes dans le Crétacé supérieur de Antunes, M.T. and Sigogneau-Russell, D. 1991. Nouvelles données sur Provence. Comptes rendus de l’Académie des Sciences de Paris, II les dinosaures du Crétacé supérieur du Portugal. Comptes rendus de 306: 153–158. l’Academie des Sciences, Paris, II 313: 113–119. Canudo, J.I. and Ruiz-Omeñaca, J.I. 2003. Los restos directos de dinosau- Antunes, M.T. and Sigogneau-Russell, D. 1992. La faune de petits dino- rios terópodos (excluyendo Aves) en España. Ciencias de la Tierra 26: saures du Crétacé terminal portugais. Comunicações dos Serviços Geo- 347–374. lógicos de Portugal 78: 49–62. Canudo, J.I., Barco, J.L., Cruzado-Caballero, P., Cuenca-Bescós, G., Ruiz- 624 ACTA PALAEONTOLOGICA POLONICA 60 (3), 2015

Omeñaca, J.I., and Royo-Torres, R. 2005. Evidencias de predación de ceous reptile bones (Laño, Iberian Peninsula): microstructural, petro- dinosaurios terópodos en el Maastrichtiense superior, Cretácico supe- logical and geochemical features. Cretaceous Research 20: 169–187. rior de Arén (Huesca). Lucas Mallada 12: 29–58. Ezcurra, M.D. and Agnolín, F.L. 2012. An abelisauroid dinosaur from the Carrano, M.T. and Sampson, S.D. 2008. The phylogeny of Ceratosauria (Di- Middle Jurassic of Laurasia and its implications on theropod palaeo- nosauria: Theropoda). Journal of Systematic Palaeontology 6: 183–236. biogeography and evolution. Proceedings of the Geologists’ Associa- Carrano, M.T., Sampson, S.C., and Forster, C.A. 2002. The osteology of tion 123 (3): 500–507. Ma siakasaurus knopfleri, a small abelisauroid (Dinosauria: Thero poda) Farlow, J.O., Brinkman, D.L., Abler, D.L., and Currie, P.J. 1991. Size, from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleon- shape and serration density of theropod dinosaur lateral teeth. Modern tology 22 (3): 510–534. Geology 16: 161–198. Chiappe, L.M., Lamb, J.P., and Ericson, P.G.P. 2002. New enantiornithine Galbrun, B., Feist, M., Colombo, F., Rocchia, R., and Tambareau, Y. 1993. bird from the marine upper Cretaceous of Alabama. Journal of Verte- Magnetostratigraphy and biostratigraphy of Cretaceous–Tertiary con- brate Paleontology 22: 170–174. tinental deposits, Ager Basin, Province of Lérida, Spain. Palaeogeog- Chiappe, L.M., Norell, M.A., and Clark, L. 1998. The skull of a relative of raphy, Palaeoclimatology, Palaeoecology 102: 41–52. the stem-group bird Mononykus. Nature 392: 275–278. Garcia, G., Duffaud, S., Feist, M., Marandat, B., Tambareau, Y., Villatte, Codrea, V., Smith, T., Dica, P., Folie, A., Garcia, G., Godefroit, P., and J., and Sigé, B. 2000. La Neuve, gisement à plantes, invertébrés et Van Itterbeeck, J. 2002. Dinosaur egg nests, mammals and other ver- vertébrés du Bégudien (Sénonien supérieur continental) du bassin tebrates from a new Maastrichtian site of the Hateg Basin (Romania). d’Aix-en-Provence. Geodiversitas 22: 325–348. Comptes Rendus Palevol 1: 173–180. Garrido Mejías, A. and Ríos Aragües, L.M.1972. Síntesis geológica del Se- Company, J. 2004. Vertebrados continentales del Cretácico superior (Cam- cundario and Terciario entre los ríos Cinca and Segre (Pirineo Central paniense–Maastrichtiense) de Valencia. 410 pp. Unpublished Ph.D. de la vertiente surpirenaica, provincias de Huesca and Lerida) Boletín Thesis, Universitat de Valencia, Valencia. Geológico and Minero 83: 1–47. Company, J., Pereda-Suberbiola, X., Ruiz-Omeñaca, J.I., and Buscalioni, Grigorescu, D. 2003. Dinosaurs of Romania: Comptes Rendus Palevol 2: A.D. 2005. A new species of Doratodon (Crocodyliformes: Ziphosu- 97–101. chia) from the Late Cretaceous of Spain. Journal of Vertebrate Pale- Huene, F. von 1914. Das natürliche System der Saurischia. Zentralblatt für ontology 25: 343–353. Mineralogie, Geologie und Paläontologie B 1914: 154–158. Cope, E.D. 1876. Descriptions of some vertebrate remains from the Fort Jagt, J.W.M., Mulder, E.W.A., Schulp, A.S., Dortangs, R.W., and Fraaije, Union beds of Montana. Proceedings of the Academy of Natural Sci- R.H.B. 2003. Dinosaurs from the Maastrichtian-type area (southeastern ences of Philadelphia 28: 248–261. Netherlands, northeastern Belgium). Comptes Rendus Palevol 2: 67–76. Cruzado-Caballero, P., Pereda-Suberbiola, X., and Ruiz-Omeñaca, J.I. 2010. Krause, D.W., Sampson, S.D., Carrano, M.T., and O’Connor, P. 2007. Blasisaurus canudoi gen. et sp. nov., a new lambeosaurine dinosaur Overview of the history of discovery, taxonomy, phylogeny, and bio- (Hadrosauridae) from the latest Cretaceous of Arén (Huesca, Spain). geography of Majungasaurus crenatissimus (Theropoda: Abelisauri- Canadian Journal of Earth Sciences 47: 1507–1517. dae) from the Late Cretaceous of Madagascar. Journal of Vertebrate Csiki, Z. and Grigorescu, D. 1998. Small theropods from the late Creta- Paleontology Memoir 8 (Supplement 2): 1–20. ceous of the Hateg Basin (Western Romania), an unexpected diversity Laurent, Y. 2002. Les faunes de vertébrés continentaux du Maastrichtien at the top of the food chain. Oryctos 1: 87–104. supérieur d’Europe. Systématique et biodiversité. Strata, Série 2 41: Csiki, Z., Vremir, M., Brusatte, S.L., and Norell, M.A. 2010. An aberrant 1–81. island-dwelling theropod dinosaur from the Late Cretaceous of Ro- Laurent, Y., Bilotte, M., and Le Loeuff, J. 2002. Late Maastrichtian con- mania. Proceedings of the National Academy of Sciences 107: 15357– tinental vertebrates from southwestern France: correlation with ma- 15361. rine fauna. Palaeogeography, Palaeoclimatology, Palaeoecology 187: Currie, P.J. 1987. Theropods of the Judith River Formation of Dinosaur Fro- 121–135. vincial Park, Alberta. In: P.J. Currie and E.H. Koster (eds.), 4th Sympo- Le Loeuff, J. 1992. Les vertébrés continentaux du Crétacé Supérieur d’Eu- sium Mesozoic Terrestrial Ecosystems. Short Papers Tyrrell Museum rope: Paléoécologie, Biostratigraphie et paléobiogéographie. 273 pp. Palaeontology 1987: 52–60. Unpublished Ph.D. Thesis, Mémories des Sciences de la Terre, Uni- Currie, P.J. 1995. New information on the anatomy and relationships of versité Pierre et Marie Curie, Paris. Dromaeosaurus albertensis (Dinosauria: Theropoda). Journal of Ver- Le Loeuff, J. and Buffetaut, E. 1991. Tarascosaurus salluvicus nov. gen., tebrate Paleontology 15: 576–591. nov., sp., dinosaure théropode du Crétacé supérieur du Sud de la France. Currie, P.J., Rigby, J.K., and Sloan, R.E. 1990. Theropod teeth from the Geobios 25: 585–594. Judith River Formation of southern Alberta, Canada. In: K. Carpenter Le Loeuff, J. and Buffetaut, E. 1998. A new dromaeosaurid theropod from and P.J. Currie (eds.), Dinosaur Systematics. Approaches and Perspec- the upper Cretaceous of southern France. Oryctos 1: 105–112. tives, 107–125. Cambridge University Press, Cambridge. Lillegraven, J.A. and Eberle, J.J. 1999. Vertebrate faunal changes through Díaz Molina, M. 1987. Sedimentación sintectónica asociada a una subida Lancian and Puercan time in Southern Wyoming. Journal of Paleon- relativa del nivel del mar durante el Cretácico Superior (Fm. Tremp, tology 73: 691–710. provincia de Lérida). Estudios Geológicos, Volumen Extraordinario Lindgren, J., Currie, P.J., Rees, J., Siverson, M., and Alwmark, C. 2009. Galve-Tremp 1987: 69–93. Theropod dinosaur teeth from the lowermost Cretaceous Rabekke For- Díaz-Molina, M., Kälin, O., Benito, M.I., Lopez-Martínez, N., and Vicens, mation on the Island of Bornholm, Denmark. Geobios 41: 253–262. E. 2007. Depositional setting and early diagenesis of the dinosaur egg- López-Martínez, N. 2003a. Dating dinosaur oodiversity: chronostratigra phic shell-bearing Aren Fm at Bastus, Late Campanian, south-central Pyre- control of Late Cretaceous oospecies succession. Palaeovertebrata 32: nees. Sedimentary Geology 199 (3): 205–221. 121–148. Díez Díaz, V., Pereda Suberbiola, X., and Sanz, J.L. 2011. Braincase anato- López-Martínez, N. 2003b. La extinción de los dinosaurios y su registro my of the titanosaurian sauropod Lirainosaurus astibiae from the Late en los Pirineos Sur-Centrales. Actas II Jornadas sobre Paleontología Cretaceous of the Iberian Peninsula. Acta Palaeontologica Polonica de Dinosaurios y su entorno, Salas de los Infantes (Burgos, September 56: 521–533. 2001), 71–98. Colectivo Arqueológico-Paleontológico de Salas, Salas Díez Díaz, V., Pereda Suberbiola, X., and Sanz, J.L. 2012. Juvenile and de los Infantes (Burgos). adult teeth of the titanosaurian dinosaur Lirainosaurus (Sauropoda) López-Martínez, N., Ardèvol, L., Arribas, M.E., Civis, J., and González from the Late Cretaceous of Iberia. Geobios 45: 265–274. Delgado, A. 1998. The geological record in non-marine environments Elorza, J., Astibia, H., Murelaga, X., and Pereda-Suberbiola, X. 1999. around the K/T boundary (Tremp Formation, Spain). Bulletin de la Francolite as a diagenetic mineral in dinosaur and other Upper Creta- Société Géologique de France 169: 11–20. TORICES ET AL.—LATE CRETACEOUS THEROPOD DINOSAURS FROM SPAIN 625

López-Martínez, N., Arribas, M.E., Robador, A., Vicens, E., and Ardèvol, Pérez-Moreno, B., Sanz, J.L., Buscalioni, A.D., Moratalla, J.J., Ortega, F., L. 2006. Los carbonatos danienses (Unidad 3) de la formación Tremp and Rasskin-Gutman, D. 1994. A unique multitoothed ornithomimosaur (Pirineos sur-centrales): paleogeografía y relación con el límite Cretáci- dinosaur from the Lower Cretaceous of Spain. Nature 370: 363–367. co–Terciario. Revista de la Sociedad Geológica de España 19: 213–255. Pol, C., Buscalioni, A.D., Carballeira, J., Francés, V., López-Martínez, N., López-Martínez, N., Canudo, J.I., Ardèvol, L., Pereda-Suberbiola, X., Orue - Marandat, B., Moratalla, J.J, Sanz, J.L., Sigé, B., and Villatte, J. 1992. -Etxebarria, X., Cuenca Bescós, G., Ruiz Omeñaca, J.I., Murelaga, X., Reptiles and mammals from the Late Cretaceous new locality Quint- and Feist, M. 2001. New dinosaur sites correlated with Upper Maas- anilla del Coco (Burgos Province, Spain). Neues Jahrbuch für Geolo- trichtian pelagic deposits in the Spanish Pyrenees: implications for the gie und Paläontologie Abhandlungen 184: 279–314. dinosaur extinction pattern in Europe. Cretaceous Research 22: 41–61. Pol., D. and Rauhut, O.W.M. 2012. A Middle Jurassic abelisaurid from Pa- López-Martínez, N., Fernández Marron, M.T., and Valle, M. F. 1999. The tagonia and the early diversification of theropod dinosaurs. Proceed- sucession of vertebrates and plants across the Cretaceous–Tertiary ings of the Royal Society of London, B 279 (1741): 3170–3175. boundary in the Tremp Formation, Ager Valley (south central Pyre- Prieto-Márquez, A., Gaete, R., Galobart, A., and Ardèvol, L. 2000. A nees, Spain). Geobios 32: 617–627. Richardoestesia-like theropod tooth from the Late Cretaceous fore- Lyson, T.R., Bercovici, A., Chester, S.G.B., Sargis, E.J., Pearson, D., and deep, south-central Pyrenees, Spain. Eclogae geologica Helvetiae 93: Joyce, W.G. 2011. Dinosaur extinction: closing the ‘3 m gap’. Biology 497–501. Letters 7: 925–928. Rauhut, O.W.M. 2002. Dinosaur teeth from the Barremian of Uña, Prov- Makovicky, P. and Norell, M.A. 2004. Troodontidae. In: D.B. Weishampel, ince of Cuenca, Spain. Cretaceous Research 23: 255–263. P. Dodson, and H. Osmólska (eds.), The Dinosauria, 2nd ed., 184–195. Rauhut, O.W.M. 2012. A reappraisal of a putative record of abelisauroid California University of California Press, Berkeley. theropod dinosaur from the Middle Jurassic of England. Proceedings Marsh, O.C. 1881. Classification of the Dinosauria. American Journal of of the Geologists’ Association 123: 779–786. Science, Series 3 23: 81–86. Rauhut, O.W.M. and Zinke, J. 1995. A description of the Barremian dinosaur Matthew, W.D. and Brown, B. 1922. The family Deinodontidae, with no- fauna from Uña with a comparison to that of Las Hoyas. In: N. Me- tice of a new genus from the Creataceous of Alberta. Bulletin of the lendez (ed.), II International Symposium on Lithographic Limestones, American Museum of Natural History 46: 367–385. Lleida-Cuenca (Spain), 9th–16th July 1995, Extended Abstracts, 123– Mey, P.H.W., Nagtegaal, P.J.C., Roberti, K.J., and Hartevelt, J.J.A. 1968. 126. Universidad Autónoma de Madrid, Cantoblanco. Lithostratigraphic subdivision of post-Hercynian deposits in the South - Reichel, M. 2010. The heterodonty of Albertosaurus sarcophagus and -Central Pyrenees, Spain. Leidse Geologische Mededelingen 41: 221– Tyrannosaurus rex: biomechanical implications inferred through 3-D 228. models. Canadian Journal of Earth Sciences 47: 1253–1261. Nagtegaal, P.J.C., Van Vliet, A., and Brouwer, J. 1983. Syntectonic coastal Riera, V., Oms, O., Gaete, R., and Galobart, A. 2009. The end-Cretaceous offlap and concurrent turbidite deposition: the Upper Cretaceous Aren dinosaur succession in Europe: The Tremp Basin record (Spain). Pa- Sandstone in the south-central Pyrenees, Spain. Sedimentary Geology laeogeography, Palaeoclimatology, Palaeoecology 283: 160–171. 34 (2): 185–218. Robador, A. 2005. El Paleoceno e Ilerdiense inferior del Pirineo occiden- Nopcsa, F. 1902. Notizen über cretacische Dinosaurier. Teil 2. Megalosau- tal: Estratigrafía y sedimentología. 456 pp. Tesis Doctoral, University rus hungaricus nov. sp. ein Theropode der Siebenburgischen Kreide. del País Vasco/EHU, Bilbao. Sitzungsberichte der Mathematisch-Naturwissenschaftlichen Classe Samman, T., Powell, G.L., Currie, P.J., and Hills, L.V. 2005. Morphometry der Kaiserlichen Akademie der Wissenschaften 3 (1): 104–107. of the teeth of western North American tyrannosaurids and its appli- Norell, M.A., Makovicky, P.J., and Clark, J.M. 2000. A new troodontid cability to quantitative classification. Acta Palaeontologica Polonica theropod from Ukhaa Tolgod, Mongolia. Journal of Vertebrate Pale- 50: 757–776. ontology 20 (1): 7–11. Sánchez-Hernández, B. and Benton, M.J. 2014. Filling the ceratosaur gap: Osmólska, H., and Barsbold, R. 1990. Troodontidae. In: D.B. Weishampel, A new ceratosaurian theropod from the Early Cretaceous of Spain. P. Dodson, and H. Osmólska (eds.), The Dinosauria, 2nd ed., 259–268. Acta Palaeontologica Polonica 59: 581–600. California University of California Press, Berkeley. Sankey, J.T. 2001. Late Campanian Southern Dinosaurs, Aguja Formation, Ösi, A., Apesteguía, S., and Kowalewski, M. 2010. Non-avian theropod di- Big Bend, Texas. Journal of Paleontology 75: 208–215. nosaurs from the early Late Cretaceous of Central Europe. Cretaceous Sankey, J.T., Brinkman, D.B., Guenther, M., and Currie, P.J. 2002. Small Research 31: 304–320. theropod and bird teeth from the Late Cretaceous (Late Campanian) Owen, R. 1842. Report on British fossil reptiles, Part ii. Report of the Brit- Judith River Group, Alberta. Journal of Paleontology 76: 51–763. ish Association for the Advancement of Science for 1841 11: 60–204. Sanz, J.L., Powell, J.E., Le Loeuff, J., Martínez, R., and Pereda Suberbiola, Pereda Suberbiola, X. 1999a. Ankylosaur dinosaur remains from the Upper X. 1999. Sauropod remains from the Upper Cretaceous of Laño (north- Cretaceous of Laño (Iberian Peninsula). Estudios del Museo de Cien- central Spain). Titanosaur phylogenetic relationships. Estudios del Mu- cias Naturales de Alava 14 (Número especial 1): 273–288. seo de Ciencias Naturales de Alava 14 (Número especial 1): 235–255. Pereda-Suberbiola, X. 1999b. Las faunas finicretácicas de dinosaurios ibéri- Seeley, H.G. 1888. The classification of the Dinosauria. Report of the Brit- cos. Zubia 17: 259–279. ish Association for the Advancement of Science 1887: 698–699. Pereda Suberbiola X. 2009. Biogeographical affinities of Late Creta- Sereno, P.C., Wilson, J.A., and Conrad, J.L. 2004. New dinosaurs link ceous continental tetrapods of Europe: a review. Bulletin de la Société southern landmasses in the mid-Cretaceous. Proceedings of the Royal géologique de France 180: 57–71. Society of London B 271 (1546): 1325–1330. Pereda Suberbiola, X. and Sanz, J.L. 1999. The ornithopod dinosaur Sheehan, P.M., Fastovsky, D.E., Barreto, C., and Hoffmann, R.G. 2000. Rhabdodon from the Upper Cretaceous of Laño. Estudios del Museo Dinosaur abundance was not declining in a “3 m gap” at the top of de Ciencias Naturales de Alava 14 (Número especial 1): 257–272. the Hell Creek Formation, Montana and North Dakota. Geology 28: Pereda-Suberbiola, X., Astibia, H., Murelaga, X., Elorza, J.J., and Gó- 523–526. mez-Alday, J.J. 2000. Taphonomy of the Late Cretaceous dinosaur - Sigé, B., Buscalioni, A.D., Duffaud, S., Gayet, M., Orth, B., Rage, J.C., -bearing beds of the Laño Quarry (Iberian Peninsula). Palaeogeogra- and Sanz, J.L. 1997. Etat des données sur le gisement crétacé supérieur phy, Palaeoclimatology, Palaeoecology 157 (3–4): 247–275. continental de Champ-Garimond (Gard, Sud de la France). Münchner Pereda-Suberbiola, X., Canudo, J.I., Cruzado-Caballero, P., Barco, J.L., Geowissenschaftliche Abhandlungen, Reihe A, Geologie und Paläon- López-Martínez, N., Oms, O., and Ruiz-Omeñaca, J.I. 2009. The last tologie 34: 111–130. hadrosaurid dinosaurs of Europe: A new lambeosaurine from the Up- Smith, J.B. 2005. Heterodonty in Tyrannosaurus rex: implications for the permost Cretaceous of Aren (Huesca, Spain). Comptes Rendus Palevol taxonomic and systematic utility of theropod dentition. Journal of Ver- 8: 559–572. tebrate Paleontology 25: 865–887. 626 ACTA PALAEONTOLOGICA POLONICA 60 (3), 2015

Smith, J.B., Vann, D.R., and Dodson, P. 2005. Dental morphology and X.-J., Sahni, A., Gomani, E.M., and Noto, C.R. 2004. Dinosaur distri- variation in theropod dinosaurs: implications for the taxonomic identi- bution. In: D.B. Weishampel, P. Dodson, and H. Osmólska (eds.), The fication of isolated teeth. The Anatomical Record Part A 285: 699–736. Dinosauria, 2nd ed., 517–606. University of California Press, Berkeley. Suzuki, S., Chiappe, L.M., Dyke, G.J., Watabe, M., Barsbold, R., and Weishampel, D.A., Csiki, Z., Benton, M.J., Grigorescu, D., and Codrea, Tsogtbaatar, K. 2002. A new specimen of Shuvuuia deserti Chiappe et V. 2010. Palaeobiogeographic relationships of the Hateg biota—Be- al., 1998 from the Mongolian Late Cretaceous with a discussion of the tween isolation and innovation. Palaeogeography, Palaeoclimatology, relationships of Alvarezsaurids to other theropod dinosaurs. Natural Palaeoecology 293: 419–437. History Museum of Los Angeles County, Contributions in Science 494: Xu, X., Clark, J.M., Mo, J., Choiniere, J., Forster, C.A., Erickson, G.M., 1–18. Hone, D.W.E., Sullivan, C., Eberth, D.A., Nesbitt, S., Zhao, Q., Her- Torices, A. 2002. Los dinosaurios terópodos del Cretácico Superior de la nandez, R., Jia, C.-K., Han, F.-L., and Guo, Y. 2009. A Jurassic cerato- Cuenca de Tremp (Pirineos Sur-Centrales, Lleida). Coloquios de Pa- saur from China helps clarify avian digital homologies (supplementary leontología 53: 139–146. information). Nature 459: 940–944. Torices, A., Ruiz-Omeñaca, J.I., Canudo, J.I., and López-Martínez, N. 2004. Xu, X. and Norell, M.A. 2004. A new troodontid dinosaur from China with Nuevos datos sobre los dinosaurios terópodos (Saurischia: Theropoda) avian-like sleeping posture. Nature 431: 838 –841. del Cretácico Superior de los Pirineos Sur Centrales (Huesca, Lleida). Zinke, J. 1998. Small theropod teeth from the Upper Jurassic coal mine of Geo-Temas 6 (5): 71–74. Guimarota (Portugal). Paläontologische Zeitschrift 72: 179–189. Turner, A.H., Pol, D., Clarke, J.A., Erickson, G.M., and Norell, M.A. 2007. Zinke, J. and Rauhut, O.W.M. 1994. Small theropods (Dinosauria, Sau- A basal dromaeosaurid and size evolution preceding avian flight. Sci- rischia) from the Upper Jurassic and Lower Cretaceous of the Iberian ence 317: 1378–1381. Peninsula. Berliner geowissenschftliche Abhandlungen 13: 163–177. Vicens, E., Ardévol, L., López-Martínez, N., and Arribas, M.E. 2001. Cor- Zanno, L.E. and Makovicky, P.J. 2013. No evidence for directional evolu- relación de alta resolución del Campaniense–Maastrichtiense, Pirineos tion of body mass in herbivorous theropod dinosaurs. Proceedings of Sud-Centrales. Geo-Temas 3 (2): 261–264. the Royal Society of London B 280: 20122526. Weishampel, D.B., Barrett, P.M., Coria, R.A., Le Loeuff, J., Xu, X., Zhao,